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Three New Species of Carcinonemertes (Nemertea, Carcinonemertidae) from the Southeastern Coast of Brazil

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Three New Species of Carcinonemertes (Nemertea, Carcinonemertidae) from the Southeastern Coast of Brazil Journal of Natural History, 2006; 40(15-16): 915-930 (^) Taylor & Francis Taylor & Francis Group Three new species of Carcinonemertes (Nemertea, Carcinonemertidae) from the southeastern coast of Brazil CYNTHIA SANTOS1, JON L. NORENBURG1 & SERGIO L. S. BUENO2 1 Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institute, Washington, DC 20560-0163, USA, and 2Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Rua do Matao, travessa 14, n° 101, 05508-900, Butantan, Sao Paulo, SP, Brazil (Accepted 31 December 2005) Abstract Three new species of Carcinonemertes from the southeastern coast of Brazil are described: Carcinonemertes divae new species, Carcinonemertes caissarum new species and Carcinonemertes sebastianensis new species. They were found, respectively, associated with the crabs Libinia spinosa, Hepatus pudibundus and Menippe nodifrons, each a newly recorded host for Carcinonemertes. Characters not previously used to describe members of the family Carcinonemertidae, such as distance from ovaries to tip of head, distance from brain to tip of head and distance from stylet to tip of head are included in the descriptions of the new species and are discussed. The locations of these new carcinonemertid worms in their respective hosts are presented in detail and a novelty regarding the infestation site is registered. Keywords: Carcinonemertidae, decapod crustacean, symbiosis, predation, Carcinonemertes divae, Carcinonemertes caissarum, Carcinonemertes sebastianensis, Brazil Introduction Nemertean worms of the family Carcinonemertidae are symbiotic egg predators of many decapod crustaceans. Owing to their life cycle, intimacy and use of chemically mediated cues from their hosts, their biology is effectively akin to parasitism; their ecological impact, however, is that of a predator because they kill individual embryos (Kuris 1993; Torchin et al. 1996). The family Carcinonemertidae comprises two genera, Carcinonemertes (Kolliker 1845), which lacks accessory stylets, and Ovicides Shields, 2001, the latter with a single species, O. juliae Shields, 2001, which has accessory stylets. Ten species of Carcinonemertes have been described so far: C. carcinophila (Kolliker 1845), with two recognized subspecies, C. c. carcinophila (Kolliker 1845) and C. c. imminuta Humes, 1942; C. epialti Coe ,1902; C. Correspondence: Cynthia Santos, Department of Invertebrate Zoology, Smithsonian Institute, PO Box 37012, National Museum of Natural History W-216, MRC 163, Washington, DC 20013-7012, USA. Email: [email protected] Published 22 August 2006 ISSN 0022-2933 print/ISSN 1464-5262 online © 2006 Taylor & Francis DOI: 10.1080/00222930600833842 916 C. Santos et al. mitsukurii Takakura, 1910; C. coei Humes, 1942; C. errans Wickham, 1978; C. regicides Shields, Wickham & Kuris, 1989; C. australiensis Campbell, Gibson & Evans, 1989; C. humesi Gibson & Jones, 1990; C. wickhami Shields & Kuris, 1990, and C. pinnotheridophila McDermott & Gibson, 1993 (Shields 2001). To date only Carcinonemertes carcinophila imminuta has been reported for Brazil, in the portunid crabs Callinectes danae Smith and Callinectes ornatus Ordway (Santos and Bueno 2001). In the present paper, three new species of the genus Carcinonemertes are described, obtained from the crabs Libinia spinosa H. Milne Edwards (Decapoda, Pisidae), Hepatus pudibundus (Herbst) (Decapoda, Hepatidae) and Menippe nodifrons Stimpson (Decapoda, Menippidae) collected on the southeastern coast of Brazil. The locations of these new worms in their respective hosts are presented in detail. The distinctive morphological characters of the new species are compared to the previously described ones; new characters are included in the descriptions and discussed. Material and methods Decapod crustaceans were collected by trawling on the southeastern coast of Brazil along the following beaches: Anchovas Beach, Sao Sebastiao Island, (23°55'S, 45°19'W); Enseada Beach, Sao Sebastiao, (23°43'S, 45°25'W), and Poco Beach, Sao Sebastiao Island, (23°45'S, 45°16'W). Crabs were also collected by hand at Figueira Beach, Sao Sebastiao (23°44'56"S, 45°24'34"W). Crabs were transported to the nearby laboratory facilities at the Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, where they were kept alive in tanks with a flow-through seawater system until dissection. Identification of crabs followed Melo (1996). The exoskeleton surface and the arthrodial membranes of the crabs were macroscopically examined for nemerteans. The dorsal carapace was removed to expose the branchial chambers. Gills and pleopods were removed with the aid of forceps and were examined under a dissecting microscope. The nemerteans were collected from crabs and placed in Petri dishes filled with seawater until the moment of taking measurements, pictures and notes on characters. Nemerteans were relaxed in a 1:1 solution of 7.5% MgCl2 (prepared with bottled drinking water) and seawater for 15-30 min, after which, length and width of body were determined with the aid of an ocular micrometer in a dissecting microscope. Measurements of internal features were made with the aid of an ocular micrometer in a compound microscope after covering the worms with a coverslip. Photomicrographs were made with a Canon PowerShot A10 digital camera. Holotype and paratypes of each new species are deposited at the Museu de Zoologia da Universidade de Sao Paulo, Brazil (Abbreviation: MZUSP). Some mucus sheaths produced by Carcinonemertes sebastianensis n. sp were prepared for scanning electron microscopy to obtain more details on their morphology. The mucus sheaths were cleaned in a 1:1 solution of 7.5% MgC12 (prepared with bottled drinking water) and seawater and were fixed in a 10% seawater formalin solution. Mucus sheaths were dehydrated in a series of graded ethanol (50, 70, 95, 100 and 100% for 10 min each), dried by the critical point method with C02 and coated with gold, then examined with a JEOL 6400 Visions scanning electron microscope. New Carcinonemertes from Brazil 917 Systematics Family Carcinonemertidae Sumner, Osburn & Cole, 1913 Diagnosis Modified by Shields et al. (1989) from Humes (1942): Monostiliferous hoplonemerteans living as symbionts (egg predators) on the gills, under the abdomen, on the apodemes, and axillae, and in or on the egg masses of decapod crustaceans. Short proboscis, reaching scarcely beyond the posterior end of the muscular portion of the esophagus. Lateral nerves lie internal to the well-developed submuscular glands. Cephalic glands well developed, with cephalic muscle fibers present. Cerebral organs lacking. Takakura's duct system present in males. Internal fertilization and oviparity occur commonly. In most species, adult worms occupy, at least temporarily, mucus sheaths secreted and attached to the setae on the pleopods and hairs of endopodites of ovigerous decapods. Embryos hatch as hoplone- mertean larvae. Genus Carcinonemertes (Kolliker, 1845) Diagnosis From Coe (1902) and amended here (italic): Nemerteans living as symbionts (eggpredators) on various species of Crustacea. Proboscis but little developed, very small in size, and extremely short, without lateral pouches of reserve stylets, but armed with central stylet and basis only; anterior proboscis very short, without distinct muscular layers, without distinct nerves, and without a thickened glandular epithelium. Cerebral organs lacking. Two ocelli. Cephalic glands massively developed. Usually oviparous, though fertilization often takes place internally. Carcinonemertes divae new species (Figure 1A-F) Diagnosis Body color varies from translucent white to orange. Two eyes, black. Anterior end of body rounded or pointed; posterior end pointed. Worms 1.3-4.3 mm long. Accessory stylets absent. Ovaries arranged in one row on each side of intestine. Takakura's duct present. Ornamented and filiform mucus sheath attached to pleopods of hosts may be present in adult worms. Material examined Fourteen females, 16 males and one larva were examined. Holotype: male, from the egg mass of Libinia spinosa; type locality: Poco Beach, Sao Sebastiao Island, Brazil, (23°45'S, 45°16'W), 18 Jul 2003; Coll. Cynthia Santos; MZUSP No. 001. Paratype: female, from the egg mass of L. spinosa; type locality: Poco Beach, Sao Sebastiao Island, Brazil (23°45'S, 45°16'W); 18 Jul 2003; Coll. Cynthia Santos; MZUSP No. 002. 918 C. Santos et al. Figure 1. Carcinonemertes divae n. sp. (A) Anterior end of female, (B) proboscis aligned in a straight line, (C) stylet and basis (arrow) in a female worm, (D) male with rounded anterior end, (E) pointed posterior end of male, with seminal vesicle, (F) adult female (arrow) on Libinia spinosa eggs. Abbreviations: b, proboscis bulb; d, diaphragm; p, posterior proboscis chamber. Etymology The species name is a noun in the genitive singular and honors Dr Diva D. Correa, a Brazilian nemertean specialist, formerly from the Universidade de Sao Paulo, who dedicated almost 40 years of her life to the study of nemerteans and greatly contributed to the knowledge of this phylum. New Carcinonemertes from Brazil 919 Description The description is based on living adults and one larva. The latter was obtained from crabs collected at Anchovas Beach. Measurements are given as mean+SE (range, number of specimens observed). Female. Body color varied from translucent white to orange; gut orange; gonads translucent white. Two eyes, black, irregular, circular or elliptical; elliptical eyes the most common shape. Found free
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